i k / 4 ) ’ COMPOSITAE 6 * NEWSLETTER Number 27 December 1995 Scientific Editor: Bertil Nordenstam Technical Editor: Gunnel Wirénius Nohlin Published and distributed by The Swedish Museum of Natural History, Department of Phanerogamic Botany, P.O. Box 50007, L | B p S-104 05 Stockholm, Sweden A Re y4 (Director: Professor Bertil Nordenstam) ISSN 0284-8422 FEB - 5 1996 NEW YORK BOTANICAL GARDEN CONTENTS Rajni Gupta & K.G. Mukerji: Studies on weeds of Delhi II - Compositae 1 J.C. Jose & P.M. Mathew: Chromosome numbers in the South Indian Heliantheae (Compositae) 7 Mesfin Tadessse, Daniel J. Crawford & E. B. Smith: New synonyms 7 in Coreopsis L. and notes on C. sect. Pseudoagarista (Compositae- Heliantheae) 11 B. Nordenstam & J. F. Pruski: Additions to Dorobaea and Talamancalia (Compositae-Senecioneae) 3] yf Comp. Newsl. 27, 1995 STUDIES ON WEEDS OF DELHI III - COMPOSITAE Rajni Gupta and K.G. Mukerji Applied Mycology Laboratory Department of Botany University of Delhi Delhi - 110007 India Abstract Various plants belonging to the family Compositae were identified among plant weeds collected from different parts of Delhi. Some of these weeds persist throughout the year, while others appear for a short duration. Introduction Noxious weeds are plants which are capable of disrupting ordinary farming operations and resisting measures for control or eradication. The uses and values of weeds have been extolled by different workers (King 1951). According to him weeds have been condemned without a fair trial. Some of the weeds pose a great threat to the field crops. Weed problems in warm regions differ from those in the temperate zone. Weed plants grow more vigorously and regenerate more quickly because of existing high temperature and higher light intensity. Perennial plantation crops, of which there are few examples in the temperate zone, generate severe perennial weed problems, particularly grasses. The nutrients which are finding their way into water courses have resulted in an explosive growth of aquatic weeds to destroy the usefulness of irrigation systems, rivers and man-made lakes. Fifteen percent of world's lands are used for grazing. In much of the tropical and sub-tropical areas pasturing is more extensive than intensive and returns are low. Depending upon the degree of competition, weeds reduce the crop yield by 10 to 15 percent. Crops differ in their ability to compete with weeds. For many diseases, weeds also serve as a collateral host. In the present investigation, some of the commonly occurring Compositae weeds of Delhi are reported. The purpose of this survey is to note and collect the facultative parasites present on the weeds, which can further be used for the control of weeds. 2 Comp. Newsl. 27, 1995 Material and Methods Regular observations of the Departmental garden and fields were made of randomly growing plants belonging to the family Compositae. Some 2-4 plants of each genus in the field and garden were regularly recorded. Data were regularly collected on the time of their appearance, reproduction, and dissemination throughout the year (Table 1). Identification of these weeds were made using standard monographs (Maheshwari 1963, Collett 1921, Bhandari 1978). Special observations were made with respect to presence of any facultative and obligate parasites on these weeds. Observations and Discussion Plants belonging to Compositae, about 10 generas, were collected from the fields and gardens of Delhi. These belonged to genera Ageratum, Bidens, Blumea, Eclipta, Galinsoga, Gnaphalium, Launaea, Sonchus, Tridax and Vernonia. Plate 1. Some of these weeds appear for a short duration and others persist throughout the year. Ageratum conyzoides is an erect annual hairy herb, 30-60 cm tall. Stems often purplish, glabrous. Leaves broadly ovate, crenate. Synflorescence of congested capitula in terminal corymbs. Heads are diseoid with purple fragrant florets. Pappus scales 5. Achenes angled, black. Ageratum conyzoides appears in the month of January, flowers in the month of February and March and seed formation takes place in the month of May. It is an abundant herb by weed on moist, sandy soils. Bidens pilosa is an erect herb, 30-100 cm tall. Leaves opposite, variable, usually consisting ot two subopposite pairs of lateral leaflets and a deeply three—lobed, terminal leaflet, the latter larger than the former, all more or less deeply lobed, glabrous, basally cuneate, petioles 3-4 cm long. Heads white or yellow, corymbosely panicled, heterogamous. Stem quadrangular, ribbed, glabrous. Pappus of 3-4 rigid, retrorsely hispid, slightly spreading, somewhat unequal awns. This weed persists throughout the year. It is commonly known as "chirchitta". Blumea bifoliata is also an erect, bushy, hairy herb, branching from base. Leaves serrate. Heads solitary on long peduncles, florets yellow. Achenes narrowly oblong, angular, pappus white. This species flowers in the month of March. Eclipta alba is an erect decumbent herb, often rooting at the lower nodes. Stem and branches sulcate, strigose with adpressed white hairs. Leaves 1-6 cm long, variable in width, usually oblong, lanceolate or elliptic, narrowed at both ends. Capitula 6-8 cm in diameter, hemispherical, solitary, unequal, axillary. Involucral bracts about 8, ovate, obtuse, strigose pubescent. Ray florets ligulate, ligules small, spreading scarcely as Comp. Newsl. 27, 1995 long as bracts, not toothed. Pappus occasionally with very minute teeth on top of cuneate achenes. Gnaphalium purpureum is a common weed during the winter months in fields and moist waste places. With the start of winter season the vegetative growth of Gna- phalium appears and it flowers in the months of January and February. After that its seed dispersal starts. It is an annual herb with 15-30 cm high, erect, simple, or branched stems. Branches ascending, usually simple, thinly lanate tomentose. Leaves 6-20 x 1-3 cm, spathulate to linear-oblong, rounded at apex. Capitula mostly in axillary clusters from upper leaves, scarcely forming a spike, 8-12 mm in diameter. Each capitulum 4mm long, sessile, campanulate. Outer involucral bracts ovate, oblong, acute and lanate, inner linear—oblong. Receptacle 1.5—2.0 mm in diameter, more or less cup-like, glabrous. Male florets 4, corolla tip puberulous, perfect florets numerous. Pappus hairs distinctly united at the extreme base, 3-4 times longer than achenes. It flowers during the month of January and dispersal of seeds takes place in the months of April and May. Launaea nudicaulis is an annual, erect, glaucous herb, with white latex and a long, thick tap root, paniculately branched, terete, sulcate, woody, more or less fistular at base. Radical leaves runcinate pinnatifid; lobes numerous, very unequal, much irregulary spinulose toothed, cauline leaves sessile, half amplexicaul, pinnatifid, or irregularly toothed. Capitula on divaricate, bracteate peduncles, 1.5 cm long, as much broad. Flowers all ligulate, ligules of peripheral florets greyish on outer surface, with 2-4 parallel, purple veins, running up to the apex, tube glandular. Achenes 6 mm long, cylindric, somewhat contracted below the apex, obscurely 3-4-toothed at base, strongly ribbed. Receptacle 4 mm in diam., concave, naked. It flowers in the month of November and December and seed formation takes place in the months of February and March. Sonchus arvensis is an erect annual 2-4 ft high. Glabrous towards the base, glandular hairy upwards. Radical leaves many, 10-20 cm long; cauline leaves sessile, oblong or obovate, entire or runcinate-pinnatifid, half amplexicaul with spreading, acute auricles, dentate, glabrous. Capitula in irregular, umbellate cymes, campanulate, 12-15 mm long and wide, pedicels glandular. Flowers yellow, involucral bracts glabrous or with a few glandular hairs, achenes 3-5 mm long. Sonchus arvensis appears in the months of October and November and flowers in the month of January, seed dispersal takes place in April and May. Galinsoga parviflora is an erect herb, stems weak, 6-18 cm, usually glabrous towards the base, roughly hairy upwards. Leaves opposite, stalked, ovate—lanceolate. Heads radiate, small, on slender stalks. Involucral bracts few, broad, green, smooth, receptacle conical, covered with small, lanceolate, toothed scales; pappus of about 15 4 Comp. Newsl. 27, 1995 spathulate, fringed scales, ligules few white, very short notched, disc florets yellow, corolla 5-toothed. Achenes angled, roughly pubescent, crowned with the pappus scales. Galinsoga persists throughout the year. Tridax procumbens is a low, straggling herb, up to 20-30 cm high. Stem hairy, leaves 2.5 cm long, 1.5—2.0 cm broad, ovate elliptic, acute, deeply inciso-dentate or pinna- tisect, clothed on both surfaces with short glandular hairs, petioles short, densely hairy. Heads 12-15 mm in diam., solitary, peduncles very long, slender, sparsely clothed with weak spreading white hairs. Outer involucral bracts very hairy, ovate, acuminate, inner bracts membranous, oblong, slightly longer than outer, pubescent on back. Ray florets ligulate, ligules yellow, deeply 3-partite. Pappus of numerous slightly unequal, feathery bristles. Achenes 2 mm long, oblong, densely silky hairy, black. Vernonia cinerea is an erect, 1.5-8 cm high, annual herb. Stem terete, ribbed, glandular, more or less greenish pubescent, branched near the apex. Leaves much variable in size and shape, upper more narrow and smaller, narrowly elliptic-ovate or lanceolate, obtuse, petioles variable. Capitula in terminal, compound corymbs, with a minute, linear bract beneath each head and with small bracts in the fork of peduncle. Involucre campanulate, 4-seriate. Pappus biseriate, white, setaceous, setae of the inner row 4-5 mm long, silky, ciliate; of outer row minute, ciliate flat. Throughout the survey only Bremia lactucae, a downey mildew, was noted on the weed Sonchus arvensis, and Puccinia butleri was recorded on Launaea nudicaulis. These pathogens are obligate parasites and hence cannot be cultured. References Bhandari, M.M. 1978. Flora of the Indian Desert. Scientific Publishers, Jodhpur. 466 pp. Collett, H. 1921. Flora simlensis. Thacker, Sping, Calcutta, 656 pp. (Reprint 1971, Dehra Dun). King, F.C. 1951. The weed problem; a new approach. Faber & Faber Ltd., London, 164 pp. Maheshwari, J.K. 1963. The Flora of Delhi. Council of Scientific & Industrial Research, New Delhi, 447 pp. poas jo jesiodsiq=q UuOnNeULIOJ psogS=S SULIDMOLA =A - MOIS OaNeIOBIA = A 70019h]0U0q DILIUII DIUOULI A suaquinzoad xopis J SISuad4sD SNYIUOS S1jnvIIpnu vavunv’y] unaandind wniyoydouy psopfiaavd vsosuyvy nqyo mda q voyofig pawn] g psopid suapig Sapiozkuo? unjosa3y P66T J2QUIPAON 0) £66] JaquIeeq{ Jo polsed ay) SuLNp pajda]]09 TY]aq JO SPI2A4 “T F1GBL Comp. Newsl. 27, 1995 Ageratum conyzoides Bidens pilosa Gnaphalium purpureum Galinsoga parviflora Sonchus arvensis Vernonia cinerea Plate 1 Comp. Newsl. 27, 1995 Comp. Newsl. 27, 1995 CHROMOSOME NUMBERS IN THE SOUTH INDIAN HELIANTHEAE (COMPOSITAE) J. C. Jose and P. M. Mathew Centre for Environment and Development Chirakkulam Road Trivandrum 695 001 Kerala India Abstract Chromosome numbers of 43 species belonging to 27 genera of the tribe Heliantheae from South India are presented. Introduction The tribe Heliantheae is well represented in South India. Hooker (1882) has reported 14 genera from this region. In addition, exotic species of a few other genera have also established, like Zinnia, Montanoa, Rudbeckia, Tithonia, Helianthus, Dahlia, Cosmos, Coreopsis etc. Karyological studies on 43 species representing 27 genera were made and the results are presented below. Materials and Methods Most of the species studied were collected from different low and high altitude regions from Kerala and Tamil Nadu in the South Indian sector of the Western Ghats. Some- times materials of the same taxon were studied from different localities. Chromosome studies were made from pollen mother cells and/or root tip cells. Chromosome preparations were made by simple acetocarmine (2%) smear and squash techniques. The voucher specimens of the collections were deposited in the herbarium of the Botany Department, University of Kerala. The results are summarized in Table 1, and the taxa are arranged alphabetically. 8 Comp. Newsl. 27, 1995 RESULTS Table 1. Chromosome Numbers in 43 species of South Indian Heliantheae (Compositae) Acanthospermum hispidum DC. Bidens biternata (Lour.) Merrill ae Source and Voucher 5011 5058 5065 Quilon ; j Peppara Ootacamund Blainvilea acmellaL.)Philipson | 17__| Trivan Avalanche 0542 E g dead? Trivan Ootacamund 5049 Ootacamund 6078 TF O Tnvandrum 5061 24 24 48 24 32 5051 g Ootacamund Ootacamund 60 — _ Trivan Trivan Bangalore Nn nN wn 2 Selam 5 Tnvan a = Trivan Yercaud Coimbatore Trivandrum Trivandrum Comp. Newsl. 27, 1995 9 so Spilanthes oleracea L Ee al Tridax procumbens L ig | Verbesina encelioides (Cav.) Benth ee gal Wedelia trilobata (Linn.) A.S. Hitchc. Wedelia calendulacea Less. Wedelia biflora C.B. Clarke Zinnia haageana Regal 22 Pen The chromosome numbers of the species reported here ranged from n = 10 — 38, and they conform to an array of basic numbers ranging from x = 9 — 19 of which x = 17 and 12 predominate. Of the 43 species studied 16 were polyploids at different levels, of which tetraploids outnumbered. Meiosis in most of the tetraploids was normal which is suggestive of their alloploid nature. However, abnormal meiosis character- ized by multivalent formation resulting in wregular anaphase separation and consequent fall in pollen fertility was noticed in one tetraploid (Wedelia trilobata, x = 15) and also in the hexaploid Bidens biternata (x = 12). Intraspecific polyploidy was observed in two species, Sigesbeckia orientalis (2x, 4x) and Synedrella nodiflora (2x, 4x). The chromosome numbers observed in most of the species agree with the counts reported from elsewhere (see Robinson et al. 1981), but in a few cases, the reports are new counts. Wedelia urticaefolia DC 10 Comp. Newsl. 27, 1995 References Fedorov, A. 1969. Chromosome numbers of flowering plants. V. L. Komarov Botanical Institute, Leningrad. Gamble, J.S. 1921. Flora of Madras Presidency, vol. II. Botanical Survey of India. Heywood, V. H., Harborne, J. B. & B. L. Turner 1977 (ed.). The Biology and Chemistry of the Compositae vol. I & II. Academic Press, London. Hooker, J. D. 1882. Flora of British India, Vol. III. Reeve and Co., London. Mathew, A. & P. M. Mathew 1987. Cytological studies on the South Indian Compositae, pp. 1-177, in P. K.K. Nair (ed.). Glimpses in Plant Research, vol. Il. Cytological Research Monographs. Today and Tomorrows Printers and Publishers, New Delhi. Robinson, H., Powell, A. M., King, R. M. & F. W. James 1981. Chromosome numbers in Compositae: Heliantheae. Smiths. Contr. Bot. 52: 1-28. Comp. Newsl. 27, 1995 NEW SYNONYMS IN COREOPSIS L. AND NOTES ON C. SECT. PSEUDOAGARISTA (COMPOSITAE-HELIANTHEAE) Mesfin Tadesse and Daniel J. Crawford Ohio State University Department of Plant Biology 1735 Neil Avenue Columbus, Ohio, 43210-1293 USA and E. B. Smith University of Arkansas Department of Plant Biological Sciences Fayetteville, Arkansas 72701 USA Abstract New synonyms in Coreopsis sections Pseudoagarista and Calliopsis are provided. Keys to the sections of Coreopsis and to the species of Coreopsis sect. Pseudoagarista are also included. The latter section is poorly understood, and the key is preliminary, aimed to stimulate further research. A lectotype for C. paludosa M.E. Jones is selected. Introduction The genus Coreopsis L. is subdivided into 11 sections (Smith 1975), ten of which are found in North America while the eleventh section, C. sect. Pseudoagarista A. Gray, is represented in Mexico and South America. The systematic relationships between the sections and the species particularly within North America have been well documented (Crawford 1970, 1976, Crawford & Smith 1983 a & b, 1985, Jansen et al. 1987, Smith 1975, 1976, 1982, 1983, 1984; as examples). There have only been few additions to the list of species from North America, north of Mexico, since Sherff's revision of the genus in 1936 and Sharsmith's work in 1938 (e.g., Smith 1974, Parker in Smith 1975). A number of new species belonging to section Pseudoagarista and 11 12 Comp. Newsl. 27, 1995 derived particularly from the Andes in Peru have, however, been published since then (Crawford 1969, Sagastegui 1969, 1970, 1982, 1988, Sagastegui & Sanchez-Vega 1971, 1981, 1989, Sanchez-Vega et al. 1994, Turner 1986). Currently about 45 species are recognized in C. sect. Pseudoagarista. This section of Coreopsis is unique in possessing taxa with invariable achene morphology but diverse foliar features. The achenes are oblanceolate to obovate with two erect, antrorsely barbed pappus awns, and numerous, long, twin-celled hairs all along the margins and often also medially on the ventral surface. The receptacular bracts (or paleae) are also unique in the genus in possessing these long, twin-celled hairs medially or all along the external surface, and also in being 2—3-cleft at the apex in some species. The paleae are detached and dispersed together with the achenes. The South American species of Coreopsis sect. Pseudoagarista have been differentiated basically on foliar morphology and pubescence of phyllaries, leaves and internodes, although some of them have also been distinguished based on capitula and ray-floret sizes. The Mexican species, which also display a highly variable leaf morphology, are shown to be cross-compatible and interfertile (Crawford 1971). Preliminary hybridization studies on seven species from South America indicated that cross-incompatibility exists between all species except for C. connata—C. senaria, and C. connata—C. obovatifolia (Smith, unpublished data). These results, besides indi- cating the significance of continued studies on cross-compatibility and interfertility of the Andean taxa, also call for further studies into the relationships between the disjunct Mexican members and the Andean plants. It is also necessary to do more field work particularly in the Andean region of South America, which is a major center of diversity of Coreopsis sect. Pseudoagarista. About 30-40 % of the species are described from the Department of Cajamarca, Province of Cajamarca in Peru (see list below). This group of Coreopsis thus offers a unique opportunity for studying speciation in isolated upland habitats if, as indicated above, many of the morphologically similar taxa happen to be cross-incompatible. Alternatively, the currently recognized taxa could represent differing populations of a few or several species situated at different altitudes and mountains with limited or no possibilities for gene exchange. Could they also be considered chromosomal or cytological races differing mainly in ploidy levels? - a phenomenon common in ferns. Chromosome numbers of n = 13, 26 and 39 are known in a few of the species (Smith et al. in press). Studies on members of this section of Coreopsis are underway, and until a more thorough treatment is provided (it may probably represent a distinct genus), the correct names for certain taxa for which the types have been studied are provided here. A key to the sections of Coreopsis is also included to portray their relationships. The species so far described in C. sect. Pseudoagarista are collated and provided with remarks or Comp. Newsl. 27, 1995 13 notes on presumed affinities. Sixteen of the species were published since the last revision of the genus. Many of the species are known from the types only or a few specimens. The authors of this paper would appreciate receiving fruits with voucher specimens of any of the species listed. Field notes on these taxa covering size of and observed variations in populations are highly sought after and solicited from botanists or collectors working particularly on the flora of Peru, Chile and Ecuador. Coreopsis sect. Pseudoagarista A. Gray Coreopsis capillacea H.B.K., Nov. Gen. Sp. 4:180 (1820); Sherff, Field Mus. Nat. Hist., Bot. 11:333 (1936). Type: Peru, Andes, Humboldt & Bonpland s.n. (P holotype, photo!). Syn. Coreopsis triloba Blake, Contr. U.S. Nat. Herb. 22: 643 (1924); Sherff, Field Mus. Nat. Hist., Bot. 11: 334 (1936), synon. nov. Type: Ecuador, near Loxa, 1865, Jameson s.n. (US holotype, K isotype!). Blake (loc. cit.) compared C. triloba with C. capillacea and stated that it "... may be distinguished by its lanceolate to lance—ovate outer phyllaries, only one-half as long as the inner, in C. capillacea lance-linear, two-thirds as long as the inner”. In the key to the species of Coreopsis, Sherff (1936:291) distinguished C. capillacea from C. triloba on foliar features, viz. ‘folia laxissime patentia vel etiam subreflexa' (for C. capillacea) and ‘folia dense adgregata, suberecta’ (for C. triloba). No other differential or diagnostic characters were cited in the description and the remarks accompanying it. Sherff (1936:334) cited the isotype of C. triloba from the herbarium at Kew (K) under C. capillacea. Close examination of specimens given under both names indicated that the alleged differences do not hold and that they all belong to one taxon. Coreopsis venusta H.B.K., Nov. Gen. Sp. 4: 180/229 (1820); Sherff, Field Mus. Nat. Hist., Bot. 11: 328 (1936). Type: Ecuador, Prov. Loja, near Loja, Humboldt & Bonpland s.n. (P holotype, photo!). Syn. Coreopsis longula Blake, Contr. U.S. Nat. Herb. 22: 642 (1924); Sherff, Field Mus. Nat. Hist., Bot. 11: 327 (1936), synon. nov. Type: Peru, Prov. Chachapoyas, 1835-1836, Mathews s.n. (GH holotype, K isotype!). Blake (loc. cit.) related C. longula to C. venusta and indicated that the differences lie in some features of the leaves, capitula, peduncle and phyllaries (Table 1). As indicated in Table 1, the characters of C. longula fall within the range of variation of C. venusta. Leaf length varies continuously between 1.5 and 7 cm, and leaf width between 0.6 and 2.1 mm. The capitula are either solitary or in groups of two or three. The peduncle length and pubescence as well as the shape and pubescence of the outer phyllaries vary quite continuously. No other distinctive features were found. Comp. Newsl. 27, 1995 14 *pojeoIput jou = { ‘snoiqe[3 = - ‘Amey = + ‘(OBeoIYD ‘AIOISTH [eIMeNY JO wUMasn Plat WO |TV) “GEL ‘Te 19 O1DORISOW = 6ELIW ‘SEZII UonoD aw uAaIN’T = CEZII'T “E87 Olmwurerer = EZ ‘OSET PSouIdsy = OYETA ‘6I vl APIS = 6IP1_ :suouttoads “AIeI[OS = *[OG ‘9]¥[OIOUR]-318AO = “OUR]-"AO ‘IROUT]-SUO[QO = “Ul]-"qo ‘a]e[OVOUe]-SUOTGO = ‘ouRT-"qo sondi]9-suo[go = *[]9-"qo :3uojqo = *[qo ‘Arey]Ayd 19no = *{[Ayd'o ‘oye[nqns-aour] = ‘qns—ue] ‘Iopenog = ‘nog :sUOnRIAIIQqGy +| + OUR] ‘12-"qo| Igo -/+) aduadsaqnd adeys *1[4yd °G “p -/+) aduadSaqnd (wid) ySuUeay apuNpeag *¢ OS Ssh F- Seee S54 i Yo A ce Sormray a susuntdeds dAQeUISIId91 pue sad4} ‘suOndLIDSap [BUISIIO UO paseq VINSuO] “Dd pue VjsNuad -~D UIIMJIq SUOSLIBdWIOD *T Ie], Comp. Newsl. 27, 1995 Coreopsis sect. Calliopsis (Reichenb.) Nutt. Coreopsis tinctoria Nutt., [var. tinctoria] Jour. Acad. Nat. Sci. Phil. 2: 114 (1821); Sherff, Field Mus. Nat. Hist., Bot. 11: 417 (1936); North Am. Fl. Ser. 2 (2): 30 (1955); Smith & Parker, Brittonia 23 (2): 168 (1971); Smith, SIDA 6 (3): 206, fig. 48-50 (1976); Cronquist, Vasc. Fl. S.E. U.S., 1: 57 (1980). Diplosastera tinctoria (Nutt.) Tausch, Hortus Canal. 16 pl. 4 (1823). Calliopsis tinctoria (Nutt.) DC., Prodr. 5: 568 (1836). Type: United States, Arkansas, Red River, Nuttall s.n. (GH holotype, BM isotype!). Syn. Coreopsis atkinsoniana (Hook.) Dougl. in Lindl. Bot. Reg. 16: Plate 1736 (1830); Sherff, North Am. Fl. Ser. 2 (2): 33 (1955). Coreopsis tinctoria Nutt. var. atkinsoniana (Dougl.) H.M. Parker in Smith, Bot. Gaz. 136 (1): 83 (1975). Calliopsis atkinsoniana Hook. Fl. Bor.-Am. 1:311 (1833), synon. nov. Type: [British] Columbia, common on the sandy banks of the Colombia from Mewries [or Menzies] Island upwards, April 1825, Douglas s.n. (K isotype!). Sherff (1955:33) maintained C. atkinsoniana Dougl. as distinct from C. tinctoria based on the winged nature of its achenes. In a biosystematic study of C. tinctoria and C. cardaminefolia DC., Smith & Parker (1971) indicated that "... the presence or absence of a wing on the achene is controlled by merely one (or two) locus (loci) ..." and redefined C. tinctoria to include taxa with winged achenes. An examination of the isotype of C. atkinsoniana revealed that the immature achenes are narrowly winged with the wings being either entire or irregularly lacerate. These resemble the F2 progeny found by Smith & Parker (1971) from the hybridization between C. tinctoria and C. cardaminefolia. Coreopsis paludosa M. E. Jones, Contr. West. Bot. 12:46 (1908); Sherff, Field Mus. Nat. Hist. Bot. 11:434 (1936), North Am. Flora, ser. 2 (2): 35 (1955). Type: Mexico, State of Chihuahua, Sierra Madre, 2100 m, 17 Sept. 1903, Jones s.n. (RSA-POM lectotype!, selected here). Syn. Coreopsis maysillesii Sherff, Brittonia 11: 188 (1959), synon. nov. Type: Mexico, Durango, 1900-2100 m, 26 road miles north of railroad at Coytes, on road to San Luis, Quebrada de San Juan, 9 Aug. 1955, Maysilles 8367 (MICH holotype, isotypes F, GH, K!, NY). Sherff (loc. cit.) described C. maysillesii as an annual herb and compared it with C. stenophylla [F. Boynton], a name currently kept under synonymy in C. tinctoria Nutt. var tinctoria (Coreopsis sect. Calliopsis; Smith 1976). The isotype of C. maysillesii at Kew has a woody base and probably represents a perennial species. Based on greenhouse grown materials which survived for two to four years, Smith (1983:551) considered C. paludosa as a perennial species. The only other specimen 16 Comp. Newsl. 27, 1995 seen of this rare species, Melchert et al. 68-20 (OS), definitely represents a perennial species. Smith (1975:80, Table 1) suggested the inclusion of C. maysillesii in C. paludosa but no formal synonymy was made in that or his other subsequent works. The achenes of the types of C. paludosa and C. maysillesii are epappose and have pectinate wings similar to those of species in C. sect. Eublepharis (Smith 1983:551, fig. 2 D and E). However, the reported base number of x = 12 for C. paludosa (Smith 1985) precludes its inclusion in that section. The phyletic relationships between sections Eublepharis and Calliopsis were studied by Smith (1983), who maintained these on differences in the basic chromosome number and the nature of the achene wings, i.e., x = 13 and achene margins pectinate for C. sect. Eublepharis; x = 12 and achene wings entire for C. sect. Calliopsis. The genetic affinities of the three species in C. sect. Calliopsis were studied by Crawford et al. (1984), who showed that C. paludosa has lowered genetic affinities to either of the species in the section. In gross vegetative morphology, C. paludosa is similar to C. falcata F. Boynton of section Eublepharis, but this has a south-east North American distribution. Thus, C. paludosa seems to connect the two sections of Coreopsis in south-eastern and south-western North America. Key to the sections of Coreopsis L. 1. Achenes and paleae densely covered with long, twin-celled, white or light brownish hairs; achenes oblanceolate to obovate with 2(-3) antrorsely barbed pappus awns, wingless; phyllaries often pubescent REPLI CTU OSG ascii sos snezit oe 2 eons Bat were os eae wae Sect. Pseudoagarista — Achenes usually glabrous or verrucose, rarely with short simple or gland-tipped multicellular hairs, or with long, twin-celled hairs along the margins; paleae glabrous; achenes obovate, oblong-elliptic, oblong to orbicular, epappose or shortly pappose, usually winged; outer phyllaries glabrous or sparsely ciliate at base or along margins; inner phyllaries usually glabrous é....:.c:0...-sereoseessssssesensnsescsenessessvessseceete panera 2 2. Leaves simple, ovate to ovate-elliptic, margins serrate, mostly 10-25 x 3.5-10.5 cm; ray-florets lemon yellow, usually 5 .........cccecceeeeee Sect. Silphidium — Leaves not with the above combined features; ray-florets yellow or orange with or without reddish/purplish blotches at base, usually 8, CAROL'S 5. c.c.ssecsochdsneciivazeoctcssabiogssdsrottvecentusxesteesutfetteanse torte teat 3 3. Ray-florets elliptic, oblong-elliptic to lanceolate, minutely 2—3-fid at the apex, concolorous and pale to bright yellOw.............s:cscsssscssescsseecseeeeeeeeceeeeeeeees 4 Comp. Newsl. 27, 1995 — Ray-florets cuneate-obovate to oblanceolate, deeply 3-4-lobed at the apex with the median lobe often bilobed and also the longest, or trullate and undulate, concolorous or with dark red/purple blotches at oe FESS CTRL ILLES 110 Ta Cc Le aaa eter eater rents UES Ae ae Ce neve | RE eee 4. Ray-florets neuter; style arm apices acute, often with decurrent sweeping hairs; paleae narrowly spathulate...............c:cscsesssseseesesseees Sect. Palmatae — Ray-florets pistillate; style arm apices cuspidate with limited sweeping hairs; paleae oblong-linear to Oblanceolate ............secescssesesessesessesesessees 5. Leaves alternate, fleshy, clustered at apices of branches; ray-florets 10-20, 20-40 mm long; erect perennial herbs of coastal California SAREE MAEACEMIE ISL ATINS occas scs ovsnicstenseds oacbvoewaste dascesereacedvenspteteaseds Sect. Tuckermannia - Leaves opposite, herbaceous to slightly fleshy, uniformly dispersed on stem or branches; ray-florets 5-7, 2-9 mm long; shrubs or Petcunial herbs of montane habitats in MEXICO ........:c600..é.csce.0esssneceseteesesoattceeeseenees 6. Leaves ovate, obovate, oblanceolate, ovate-lanceolate, or oblong- ovate, serrate; receptacle flat; tube of disc-floret corolla pubescent on TET anh ee er neeenen epamianen tr esr We er rire ts ee EP Sect. Electra — Leaves or leaf segments linear to linear-filiform, entire; receptacle conic; tube of disc-floret corolla glabrous ............ccseccseseeeeeeeeenees Sect. Anathysana 7. Leaves alternate or crowded at the stem base; capitula solitary; ray- florets pistillate, truncate to shallowly 3-lobed; disc-floret corolla constricted in middle and pubescent; paleae oblong-lanceolate............seseeseeeeeeees — Leaves opposite; capitula cymose or corymbose; ray-florets sterile, deeply 3-4-lobed; disc-floret corolla not constricted and pubescent in middle; paleae narrowly linear with attenuate or spathulate apeX.............scsceseseeee 8. Disc-floret achenes oblanceolate, glabrous or with short, tubercle- Based OF plandular Hairs, CXALISIALE ............ccsseesssectecsssecsosooseacssonevacess Sect. Leptosyne — Disc-floret achenes oblong-elliptic with long, twin-celled hairs along the margins; pappus awns 2, paleaceous, flat, antrorsely hispid No 20 co n0 5 dnussuctentcatens Vases vicnassoncdevostoentedeanas setee crete Sect. Pugiopappus 9. Disc-floret corolla 5-lobed; leaves OPPOSILE ............:eseecseseeseeteeees Sect. Coreopsis — Disc-floret corolla 4-lobed; leaves alternate OF OPPOSILE .............seceeeeeeeeeeeeneeees 10. Achenes oblong-elliptic, outer surface rounded, inner surface flat, wings entire to pectinate; leaves opposite (rarely alternate), simple to pinnatifid; paleae linear-attenuate (X = 12)... eeeesetceeeeeeeeeeeee Sect. Calliopsis 18 Comp. Newsl. 27, 1995 — Achenes oblong, flattish, wings deeply fimbriate-lacerate; leaves simple to auricled; paleae linear, oblanceolate to spathulate (x = 13) RON ois ef Een Lee ShR CR Ce ee Bt Pare RE RAR BROS Sect. Eublepharis Taxa in C. sect. Pseudoagarista A. Gray C. breviligulata Sagast. & Sanchez (1981). Type: Peru, Dept. Cajamarca, Prov. Cajamarca, between San Marcos and Matara (bajada a Guayobamba), 2400 m, 6 May 1973, Sanchez Vega 1130 (HUT holotype, Herb. Cazamarques, isotype). Sagastegui & Sanchez (1981:225) compared it with C. peruviana. The differences lie in texture of leaves, size of capitula and shape of ray-florets. C. cajamarcana Sagast. & Sanchez (1989). Type: Peru, Dept. Cajamarca, Pullucana (Banos del Inca-La Encanada), 2700 m, 17 June 1975, Sagastegui, Cabanillas & O. Dios 8064 (Herbario de la Universidad Anterior Orrego de Trujillo, holotype). Sagastegui & Sanchez (1989: 24) related this to C. senaria. The differences lie in the achenes, size of leaves, and phyllaries. C. canescentifolia Sagast. (1988). Type: Peru, Dept. Cajamarca, Prov. Celendin, La Tranca (alrededores de Celendin), 2700 m, 21 May 1976, Sagastegui, Cabanillas & O. Dios 8498 (HUT holotype). Sagastegui (1988:51) related this to C. holodasya and C. pervelutina. The similarities are in leaf type, i.e. entire or simple, and the differ- ences lie in the shape of the upper leaves and their pubescence. C. capillacea H.B.K. (1820). Type: ?Peru, Andes, Humboldt & Bonpland (P holo- type). Compare with C. fasciculata and C. townsendii. C. celendinensis Sagast. & Sanchez (1971). Type: Peru, Dept. Cajamarca, Prov. Celendin, La Chocta—Oxamarca road, 3300 m, 30 July 1970, Sanchez s.n. (HUT holotype, Caxamarques, LP, isotypes). Sagastegui & Sanchez (1971:340) compared it with C. connata and listed a number of differences in leaf shape, dentation, capitula size, number of nerves on outer phyllaries and paleae and shape of ray-florets. C. connata Cabrera (1962). Type: Peru, Dept. Cajamarca, Prov. Celendin, La Tranca, 2600 m, 22 May 1960, Lopez & Sagastegui 3124 (LP holotype). Compare with C. helleborifolia. C. dentifolia Sanchez, Sagast. & Crawford (1994). Type: Peru, Dept. Cajamarca, Prov. Chota, District Paccha, c. 1 km E of Paccha, 2000 m, 21 July 1993, Sanchez V. 6559 (CPUN holotype, F, HAO, MO, OS, isotypes). Sanchez et al. (1994:39) stated that it is different from the other Peruvian species on account of its petiolate leaves with ovate lamina and related it to C. canescentifolia and C. pervelutina (but compare with C. woytkowskil). Comp. Newsl. 27, 1995 C. dilloniana Sanchez, Sagast. & Crawford (1994). Type: Peru, Dept. Cajamarca, Prov. Hualgayoc, 5-6 km S. of Bambamarca, c. 2780 m, 22 April 1993, Dillon & Sanchez V. 6471 (F holotype, CPUN, HAO, OS, isotypes). Sanchez et al. (1994:42) related it to C. lopez-mirandae, but compare also with C. woytkowskii. C. fasciculata Wedd. (1855). Type: Peru, Gay s.n. (P holotype). Compare with C. pickeringii and C. townsendit. C. fasciculata Wedd. var laevigata Sherff (1935). Type: Peru, Cuzco, 3400-3600 m, Herrera 111 (B holotype). Compare with C. notha var parvulifolia. C. ferreyrae Sagast. & Sanchez (1989). Type: Peru, Dept. Cajamarca, Prov. Celendin, Hacienda Limon, 2150 m, 5 May 1970, Sagastegui 7416 (HUT holotype, CPUN, F, isotypes). A species noted for its much dissected leaves with xeromorphic aspect. C. foliosa A. Gray (1861). Type: Peru, Taulia, 1835, Mathews 1376 (K holotype). Compare with C. nodosa. C. glaucodes Blake & Sherff ex Sherff (1925). Type: Peru, Dept. Cajamarca, below Hacienda La Tahona, 2600 m, near Hualgayoc, 15 May 1904, Weberbauer 4048 (B holotype, US isotype). C. helleborifolia Sanchez, Sagast. & Crawford (1994). Type: Peru, Dept. Cajamarca, Prov. San Marcos, between Chancay and Condebamba valley, 2600 m, 26 March 1994, Sanchez & Sanchez 6923 (CPUN holotype, HAO, HUT, F, MO, OS, isotypes). Sanchez et al. (1994:44) related it to C. breviligulata, but compare with C. connata. C. holodasya Blake ex Parker (1962). Type: Peru, Dept. Apurimac, Saihuiti, 4270 m, 14 May 1939, Balls 6846 (K holotype, US isotype). C. imbricata Sherff (1930). Type: Peru, Cordillera Huante, between Lima and Cuzco, 3600 m, Feb. 1867, Pearce s.n. (K holotype). Sherff (1930:371) related this to C. pickeringii and noted that "... it differs in its numerous, much shorter internodes, its lack of elongate peduncles, the caudate-tipped stigmas of its disk florets, its double outer involucre the bracts of which are larger (with a most noticeable imbricate appearance), etc." C. integra Blake (1937). Type: Peru, Dept. Huanuco, Sawn Carlos Mines, 6 miles west of Huallanca, 2745 m, 30 Sept. 1922, Macbride & Featherstone 2469 (F holotype, US isotype). Blake (1937:386) compared it with C. longula which is now a synonym of C. venusta. C. irmscheriana Bruns (1929). Type: Peru, Lomas, Mejia, Ravine of Chule, 30 m, 12 Nov. 1923, Guenther & Buchtien 55 (HBG holotype, M isotype). This is the only annual species in this section, and it is so far known only from the type specimen; note the altitude; more specimens needed. 20 Comp. Newsl. 27, 1995 C. killipii Sherff (1933). Type: Peru, Dept. Junin, Tarma, 3000-3200 m, 20-22 April 1929, Killip & Smith 21823 (F holotype). Sherff (1933:595) related this to C. macbridei and C. notha and stated that it differs from the former "... in having smaller leaves, fewer and much larger heads, and also comparatively (and actually) much larger exterior involucral bracts, etc." and from the latter "... it differs in having smaller leaves, linear-oblong (not lanceolate to oblong) leaf segments, about 8 (not 6) exterior involucral bracts, these 3—5 (not 2-3) mm long, etc." C. lopez-mirandae Sagast. (1969). Type: Peru, Dept. Amazonas, Prov. Chachapoyas, between Paso de Calla-Calla and Balsas, 3000 m, 1 June 1963, Lopez, Sagastegui & Collantes 4460 (HUT holotype, LP isotype). Sagastegui (1969:250) compared it with C. woytkowskii and noted differences in leaf margins and shape, inflorescence type, shape, nerve number and apical features of outer phyllaries and apex of paleae. C. macbridei Sherff (1930). Type: Peru, Huanuco, 2100 m, 26 April 1923, Macbride 3504 (F holotype, US isotype). Compare with C. townsendii. C. mcvaughii Crawford (1969). Type: Mexico, Aguascalientes, Sierra del Laurel, c. 10 m. SE of Calvillo, McVaugh & Koelz 225 (MICH holotype, MICH isotype). C. microlepis Blake & Sherff ex Sherff (1925). Type: Peru, Dept. Amazonas, Prov. Chachapoyas, 1835-1836, Mathews 1418 (K holotype, US isotype). Sherff (1925:372) related this to C. foliosa, C. glaucodes and C. parviceps. C. nodosa (1933). Type: Peru, Dept. Cuzco, Ollantaitambo, 2900-3100 m, 26 April 1925, Pennell 13646 (F holotype). C. notha Blake & Sherff ex Sherff (1925). Type: Peru, Dept. Cajamarca, Prov. Cajamarca, above San Pablo, 2500-2700 m, 26 April 1904, Weberbauer 3812 (B holotype, US isotype). Sherff (1925: 373) wrote that it is near C. spectabilis, but compare with C. foliosa. C. notha Blake & Sherff ex Sherff var. parvulifolia Sherff (1935). Type: Peru, Dept. Cajamarca, Prov. Contumaza, Contumaza, 1875, Raimondi 6695 (B holotype). Sherff (1935:707) noted this to be similar to C. fasciculata var. laevigata. C. oblanceolata Blake (1924). Type: Peru, western Cordillera, opposite Huan- cabamba, 2400-2850 m, 26 Sept. 1911, Townsend A211 (F holotype). C. obovatifolia Sagast. (1982). Type: Peru, Dept. Cajamarca, Prov. Cajamarca, La Encanada, 2600 m, 17 Aug. 1973, Sagastegui 7771 (HUT holotype). Sagastegui (1982:265) related this to C. oblanceolata. Comp. Newsl. 27, 1995 21 C. parviceps Blake & Sherff ex Sherff (1925). Type: Peru, Tambillo, 19 Aug. 1878, De Jelski 765 (B holotype, US isotype). Sherff (1925:369) related this to C. foliosa and C. glaucodes and stated that it differs in leaf dissection and pubescence, and shape and pubescence of the outer phyllaries. C. peruviana Sagast. (1970). Type: Peru, Dept. Cajamarca, Prov. Celendin, 3050 m, 8 May 1970, Sagastegui 7495 (HUT holotype, Herb. Caxamarques, isotype). C. pervelutina Sagast. (1982). Type: Peru, Dept. Cajamarca, Prov. Bambamarca, between Bambamarca and Hualgayoc, 2900 m, 7 Aug. 1973, Sanchez V. 1198 (HUT holotype, Herb. Caxamarques, isotype). Sagastegui (1982:267) related it to C. holodasya, but compare also with C. canescentifolia. C. petrophila A. Gray ex S. Watson (1887). Type: Mexico, Jalisco, Rio Blanco, Palmer 530 (US holotype, GH, MO, NY, UC, US, isotypes). C. petrophiloides Robinson & Greenman (1894). Type: Mexico, Jalisco, Nevado de Colima, Pringle 5508 (GH holotype). C. pickeringii A. Gray (1861). Type: Peru, Dept. Lima, above Osrajillo, U.S. Explor- ing Expedition under Captain Wilkes s.n. (US holotype). Compare with C. townsendit. C. piurana Sherff (1964). Type: Peru, Dept. Piura, near Huancabamba, May 1958, Soukup H662 (US holotype). Sherff (1964:371) traced this to C. pickeringii in his key to Coreopsis (Sherff 1936) and stated that the two plants are different. Unfortunately the most important differences in many members of this section, ie., leaf characteristics, are omitted from the description. C. polyactis Blake & Sherff ex Sherff (1925). Type: Peru, Dept. La Libertad, Prov. Otuzco, 3500-3600 m, between hacienda Llaguess and Succhabamba, 28 June 1914, Weberbauer 6995 (B holotype, US isotype). C. pringlei Robinson (1907). Type: Mexico, Queretaro, San Juan del Rio, Pringle 10050 (GH holotype, F, MICH, MO, NY, US, isotypes). C. queretarensis B.L. Turner (1986). Type: Mexico, Queretaro, Mpio. Penamiller, Maguey Verde, 17 km SE of Pinal de Amoles, 2170 m, 22 Oct. 1982, Tenorio & Romero 2353 (TEX holotype, MEXU isotype). C. rhyacophila Greenman (1900). Type: Mexico, Morelos, near Cuernavaca, Pringle 7866 (GH holotype). C. rudis (Bentham) Hemsley (1881). Type: Mexico, Jalisco, Bolanos, Canton of Colotan, 1836-1838, Hartweg 116 (K holotype, B, GH, K, isotypes). C. senaria Blake & Sherff ex Sherff (1925). Type: Peru, Pacasmayo to Moyobamba, April-June 1875, Stubel 35 p.p. (B holotype, US isotype). 22 Comp. Newsl. 27, 1995 C. sherffii Blake (1937). Type: Peru, Dept. Huanuco, Chinchapalca, 6 miles above Mito, 2900 m, 16-27 July 1922, Macbride & Featherstone 1596 (F holotype, US isotype). Blake (1937:387) compared it with C. microlepis. C. spectabilis A. Gray (1861). Type: Peru, Andes, McLean s.n. (K holotype). C. suaveolens Sherff (1930). Type: Chile, Dept. Tarapaca, Prov. Tarapaca, Cordillera de Lallinca, 3800 m, March 1926, Werdermann 1114 (GH holotype, F, S, UC, isotypes). Sherff (1930:369) related this to C. fasciculata. C. suaveolens Sherff var. ecuadoriensis Sherff (1964). Type: Ecuador, Prov. Azuay, Paramos de Silvan, Nudo de Cordillera occidental y Cordillera oriental, 3000 m, 30 July—3 Aug. 1959, Barclay & Juajibioy 8563 (US holotype). C. townsendii Blake (1924). Type: Peru, Huascaray, 1950-2250 m, 10 Sept. 1911, Townsend A 192 (F holotype). C. venusta H.B.K. (1820). Type: Ecuador, Prov. Loja, near Loja, Humboldt & Bonpland s.n. (P holotype, B isotype). C woytkowskii Sherff (1936). Type: Peru, Dept. Cajamarca, Prov. Celendin, 2625 m, 5 June 1936, Woytkowski 24 (F holotype). Acknowledgements Research grant from the National Science Foundation (No. INT- 93 155 27) is gratefully acknowledged. We thank the directors and curators of the herbaria at F, HGB, K, MO, OS, P and WAG for the loan of specimens. References Blake, S. F. 1924. New American Asteraceae. Contrib. U.S. Nat. Herb. 22: 587-661. Crawford, D. J. 1969. A new species of Coreopsis (Compositae) from Mexico. Brittonia 21 (4): 353-354. Crawford, D. J. 1970. Systematic studies on Mexican Coreopsis (Compositae). Coreopsis mutica: flavonoid chemistry, chromosome numbers, morphology, and hybridization. Brittonia 22: 93-111. Crawford, D. J. 1971. Systematics of the Coreopsis petrophiloides-lucida-teotepensis complex. Amer. J. Bot. 58 (4): 361-367. Comp. Newsl. 27, 1995 23 Crawford, D. J. 1976. Taxonomy of Coreopsis sect. Pseudoagarista (Compositae) in Mexico with additional comments on sectional relationships in Mexican Coreopsis. Brittonia 28: 329-336. Crawford, D. J. & E. B. Smith 1983a. Leaf flavonoid chemistry of North American Coreopsis (Compositae): intra- and intersectional variation. Bot. Gaz. 144 (4): 577- 583. Crawford, D. J. & E.B. Smith 1983b. The distribution of anthochlor floral pigments in North American Coreopsis (Compositae): taxonomic and_ phyletic interpretations. Amer. J. Bot. 70 (3): 355-362. Crawford, D. J. , Smith, E. B. & R. E. Pilatowski 1984. Isozymes of Coreopsis section Calliopsis (Compositae): genetic variation within and divergence among the species. Brittonia 36 (4): 375-381. Crawford, D. J. & E.B. Smith 1985. Leaf flavonoid chemistry and taxonomy of Coreopsis sect. Coreopsis. Biochemical Systematics and Ecology 13 (2): 115-118. Jansen, R. K., Smith, E. B. & D. J. Crawford 1987. A cladistic study of North American Coreopsis (Asteraceae: Heliantheae). Pl. Syst. Evol. 157: 73-84. Sagastegui, A. A. 1969. Cuatro especies nuevas de Compuestas Peruanas. Bol. Soc. Arg. Bot. 11 (4): 240-250. Sagastegui, A. A. 1970. Tres Compuestas Austroamericanas nuevas 0 criticas. Bol. Soc. Bot. La Libertad 2: 73-75. Sagastegui, A. A. 1988. Dos nuevas especies de Compuestas Peruanas. Bol. Soc. Bot. La Libertad 14: 49-53. Sagastegui, A. A. & I. Sanchez Vega 1971. Una nueva especie de Coreopsis (Compositae) del Peru. Bol. Soc. Arg. Bot. 13: 337-340. Sagastegui, A. A. & I. Sanchez Vega 1981. Un nuevo taxon del genero Coreopsis (Compositae). Darwiniana 23 (1): 223-225. Sagastegui, A.A. & I. Sanchez Vega 1989. Novedades en el genero Coreopsis. Bol. Soc. Bot. La Libertad 15 (1): 23-28. Sanchez Vega, I., Sagastegui, A. A. & D. J. Crawford 1994. Compuestas Andino- Peruanas nuevas para la ciencia. V. Arnoldoa 2 (2): 37-45. Sharsmith, H. K. 1938. The native Californian species of the genus Coreopsis L. Madrono 4: 209-231. Sherff, E. E. 1925. New or otherwise noteworthy Compositae, II. Bot. Gaz. 80 (4): 367-389. 24 Comp. Newsl. 27, 1995 Sherff, E. E. 1930. New or otherwise noteworthy Compositae, IV. Bot. Gaz 89 (4): 362-373. Sherff, E. E. 1933. New or otherwise noteworthy Compositae, VIII. Bot. Gaz. 94 (3): 589-597. Sherff, E. E. 1935. New or otherwise noteworthy Compositae, X. Amer. J. Bot. 22: 705-710. Sherff, E. E. 1936. Revision of the genus Coreopsis. Field Mus. Nat. Hist. (Bot.), 11(6): 279-475. Smith, E. B. 1974. Coreopsis nuecensis (Compositae) and a related new species from southern Texas. Brittonia 26: 161-171. Smith, E. B. 1975. The chromosome numbers of North American Coreopsis with phyletic interpretations. Bot. Gaz. 136 (1): 78-86. Smith, E. B. 1976. A biosystematic survey of Coreopsis in eastern United States and Canada. SIDA 6 (3): 123-215. Smith, E. B. 1982. Phyletic trends in section Coreopsis of the genus Coreopsis (Compositae). Bot. Gaz. 143 (1): 121-124. Smith, E. B. 1983. Phyletic trends in sections Eublepharis and Calliopsis of the genus Coreopsis (Compositae). Amer. J. Bot. 70: 549-554. Smith, E. B. 1984. Biosystematic study and typification of the Californian Coreopsis (Compositae) sections Tuckermannia, Pugiopappus, and Euleptosyne. SIDA 10 (4): 276-289. Smith, E. B. & H. M. Parker 1971. A biosystematic study of Coreopsis tinctoria and C. cardaminefolia (Compositae). Brittonia 23 (2): 161-170. Smith, E. B., Sagastegui, A., Sanchez Vega, I. & D. J. Crawford (in press). Chromosome numbers in Peruvian Coreopsis (Asteraceae: Coreopsideae). Arnoldoa. Turner, B. L. 1986. A new species of Coreopsis sect. Pseudoagarista (Asteraceae) from Mexico. Brittonia 38 (2): 168-170. Comp. Newsl. 27, 1995 Appendix A preliminary key to Coreopsis sect. Pseudoagarista A. Gray . Annual herbs; leaves simple, ovate to ovate-lanceolate, serrate ......C. irmscheriana Shrubs, subshrubs or perennial herbs; leaves simple to thrice famamatiscet, marpins entire to COAarSely SCITALE:......:...:.ccssssecessssssorsenesescassneoeazcesneedten? 2 Pebcaves ample; Margins entire Or SEmrates.s..:26s.eekiee sede RTAU i eeanecccteneccueasnse 3 Pere ces NOME. (CO LriCE-PINNAL SEC baii.idenc. 265 eee eee. acne weceserescesnutteeeee ete 1) Me AEA OUNS SCITALE £0) INCISEM «.....ncccs.c4 tacos decees ea tee et eceen cesectereonecteetonctee eee cee eae 4 Leaf margins entire, sometimes thickened and inflexed or purple MM ooo acnacinn tecsacbst aod ones deohiassevcees Ree ae ES Ee 8 . Leaves rounded or orbicular to ovate, sessile, margins regularly eoimeacua erossly, Serrate-lOWEd.).asscccs.-.2edoreess cteeteeean cee cb teeter e codec nee 5 Leaves oblanceolate, obovate, linear, linear-elliptic, subulate or Ranretece ee NTNU CLES. vs strostonosch'an entvn' osebdeonnen cece yeeateeo tear heheat nets ae ee ieee hws eee 6 . Capitula solitary or 1-3 per branch; ray-florets 8-10 x 4.4-6 mm; outer phyllaries obovate-spathulate, about equal to the inner ones. .......... C. connata Capitula 3-18, cymosely arranged, ray-florets 3-3.5 x 22.5 mm; outer phyllaries oblong-linear, 1/3 to 1/2 of the inner ones............. C. helleborifolia . Ray-florets 8.5—10 x 5—5.5 mm; capitula 28-32 mm wide at anthesis.. C. dentifolia Ray-florets 14-25 x 7-10 mm; capitula 30-50 mm at anthesis ............::cceceeeeeeees 7 . Leaves ovate-elliptic to oblanceolate, margins regularly serrate or serrate only near the apex; outer phyllaries densely pubescent on both surfaces; inner phyllaries pubescent on outer surface; achenes 6-10 ROAR EERHOIN DRS Pio eee wc kec Sk bav sade sein svebeevooe de sotto stoarontey stescatens tose demetn oateete C. woytkowskii Leaves oblanceolate, margins entire, apex 3—5-lobed or incised; phyllaries glabrous except for the ciliate margins on outer ones; PRBS G SPS) MND LONG acc sa cssc ses sanserdSoesoctsrncee ces csecett caocteanesattese C. lopez-mirandae alecavesiGensely appressed LOmeNtOSe,, Bray QTC s.,...25c0s-0--e8e0-s000se eee ae 33 32. Hairs on internodes and peduncle dense, eglandular; phyllaries usually pubescent at least at base or on margins; terminal leaflets AG Trine LONG 5, inch e- carey ncvsseunls Boner eeeko vee sess weskonddaden asta C. foliosa - Hairs on internodes and peduncle sparse, short, capitate-glandular; phyllaries glabrous; terminal leaflets 2-4 mm long (usually whole leaf and leaflets much smaller than in C. fOli0SA)........ccccccccsccssssesseseeseeeeeee C. nodosa 33. Peduncle and/or phyllaries sparsely to densely pilOse .............:.scscesceseseeseeeeseeeseeees 34 - Peduncle and phyllaries glabrous or glabrescent ..............:cscsscssessscerseceserecersececees 48 34. Both peduncle and phiyllanies pilose ...:.::c0ossssrecsecseososvessososenennsureosedresidecs Nate 35 - Atleast outer or inner phyllaries or both glabrous..............scscsscessessceeerssrcencereeees 44 Comp. Newsl. 27, 1995 35. Capitula 35-50 x 9-11 mm wide at anthesis; inner phyllaries MMMM ROP IANO NOIR ios sacs seas sau csahagévs sates ihsailses=x sence cushseeacauteceeett on ses ates ta ae 36 - Capitula 12-34 (—38) x 5~9 (-10) mm at anthesis; inner phyllaries eae MINE (OMAP SoS cde este cat vctn ct tases ces Yancs Sh Sra eewiateacEedaaene me reeaee tee 39 36. Peduncle S—10 mm long; outer phyllaries 6-8 mm long; leaf amb MARGE) eC) SCOIITANA WECLE ceo 00 soSSae ate ceva cc cere cattaenoscssececoaceese@rscecocetaecetonresee eee 37 - Peduncle 50-150 (230) mm long: outer phyllaries 4-6 mm long; Ei EESUTGT 10 og Dio 00) 1 2 (6 oie onan ee nice fo 2 ie 38 37. Leaves pinnately 3—5-lobed, ciliate on margins, all exceeding internodes; outer phyllaries broadly ovate-lanceolate, 5—7-veined, Sd. TL TLCS ee cee eee ee RO te ae C. polyactis - Leaves ternately bipinnatisect, glabrous except for tufts of hairs at petiole bases; outer phyllaries oblong, 3—S-veined, 1.1—1.6 mm wide 1 COPED EEE Eee oT EE Arcee eet ERE be C. imbricata 38. Inner phyllaries pilose at least at base; capitula 35—SO mm wide at pmbsesiss FeAVeES 2035 mii JONG ac. esedsasizeisesesssdasse denoesenatdovaccssessepatés C. pickeringii - Inner phyllaries glabrous; capitula 12-38 mm wide at anthesis; leaves NES MBERTERUDL Oe Se Soe de oc soda ccsogais ce docoen co A esos dasancad ed tae Sonos C. capillacea 39. Ray-floret lamina 8-9 X 5.5-6.5 mm; involucre 5—11 X S—7 mm at STU TS accede eee eae on Oe no me ore C. dilloniana - Ray-floret lamina 14-17 x 5-8 mm; involucre c. 10 x 12 mm at SOU sO ca csc recat wes hus Pion aan cbse dnce Mavs be vesazs testucat Sheses daeateare Goer 40 40. Capitula 5-7 mm wide at anthesis; outer phyllaries 2.54.5 mm long.................. 4] - Capitula 7-11 mm wide at anthesis; outer phyllaries 4-7 mm long...................5 42 41. Petiole S—10 mm long; outer phyllaries glabrous or pubescent; inner ols EGRESS 57 7210) (01) ISAS On Oe = nae De Re ea C. fasciculata - Petiole 10-25 mm long; outer phyllaries sparsely pubescent at base; ine TPAYUATICS SPATSCly PUDESCENL. ..:2..c..sar-cere.dececaeecenendneeeseoncnducseattes C. townsendii 42. Leaves 30-60 mm long; peduncle 50-100 mm long; inner phyllaries ST TDS coco Rae ee MEAS C. capillacea - Leaves 12-25 mm long; peduncle 20-30 (50) mm long; inner PARE ABA SUTIN OSC as o'r 'sa ago iaseacs seasasabgisevsaeesdects teense ee ee ee 43 29 30 Comp. Newsl. 27, 1995 43. Ray-floret lamina 7-11 mm long; capitula up to 23 mm wide at SERAETMSES Get Le, aeee ob Raca ca cl Seteh adil Wheel Madtnvanteanerecansanenemnteteareatee C. suaveolens - Ray-floret lamina c. 15 mm long; capitula 25—30 mm wide.................. C. piurana A4. Ray-floret lamina’7.5—14 X 3=6:5 MANN .v...ccecscsacsesseseosccscesccssctesccastecsuteonaeseeeeeeeeee 45 - jRay-floret lamina’l5—23°X-7—10.5 mim..............sece.sscenstvecessenstaeanceqeeeeee one 46 45. Leaf segments 2-6 mm wide; outer phyllaries 1.5—2 mm long; Fay-floret TAMINa T= % S35 MIN 5... Sodecckesssssedes sateadis aap oneseh hsveneeoeee C. microlepis - Leaf segments 0.3—-1 mm wide; outer phyllaries 2.5-4.7 mm long; Pay Mre tS NOTA RSG 5 WI ssn spenasnncazsoneeneWatas cudese tenet eecavo erate C. capillacea 46. Capitula 5-7 mm wide at anthesis; petiole S—10 mm long.................. C. fasciculata - Capitula 8-13 mm wide at anthesis; petiole 10-20 mm Ion... eeeeseeneeseeee 47 47. Outer phyllaries glabrous; leaf segments 1-3 mm Wide.............seeeeseneeeenees C. notha - Outer phyllaries pubescent; leaf segments 0.5—1 mm wide................ C. capillacea 48. Peduncle swollen near base of receptacle, 220-270 mm long; capitula 35-40 mm wide at anthesis; ray-floret lamina 15—18 x 10 mm........... C. spectabilis - Peduncle uniform throughout, 10-120 mm long; capitula 20-28 mm wide at anthesis; ray-floret lamina 8-12 X 1.5—7 MM...........cscsseseeeeeeeeneeneeeneens 49 49. Leaves bipinnate with the terminal segment equal to the laterals; involucre glabrous; outer phyllaries triangular ..............ceeeseeeseeseeeeeeees C. ferreyrae - Leaves pinnate with the terminal segment equal to or longer than the laterals; involucre bearded or sparsely pilose; outer phyllaries oblong- BUCA iceiacisvn azvudavitehsnsysnsddagnysovelaussaslecdintessduviesncistssanes Qovsiztataidesssis aoteee ee eee 50 50. Capitula several to numerous, in corymbose cymes; peduncle 1.5— 5.5 cm long; terminal leaf segment much longer than the laterals......... C. macbridei - Capitula solitary or in 2's or 3's; peduncle up to 12 cm long; terminal leaf segment equal to or only slightly longer than the laterals.................+ C. killipii Comp. Newsl. 27, 1995 31 ADDITIONS TO DOROBAEA AND TALAMANCALIA (COMPOSITAE-SENECIONEAE) B. Nordenstam ) and J. F. Pruski 2) 1) Department of Phanerogamic Botany Swedish Museum of Natural History P. O. Box 50007 S-104 05 Stockholm, Sweden 2) Department of Botany, MRC-166 National Museum of Natural History Smithsonian Institution Washington, D.C. 20560, USA Abstract Three new combinations are published, viz. Dorobaea laciniata (H.B.K.) B. Nord. & Pruski, D. callacallensis (Cuatr.) B. Nord. & Pruski, and Talamancalia putcalensis (Hieron.) B. Nord. & Pruski, raising to three, the number of species in each genus (both of the Compositae-Senecioneae). Talamancalia is a new generic record for South America, and was previously known only from Panama and Costa Rica. Keys to the species of Dorobaea and Talamancalia are provided. Dorobaea Cass. In 1818 Humboldt, Bonpland, and Kunth described Senecio pimpinellifolius and three other closely related species (S. nubigenus H.B.K., S. pedicularifolius H.B.K., S. lac- iniatus H.B.K.), all from the Ecuadorian Andes (Humboldt et al. 1818). The authors were struck by the distinctive habit of these taxa and suggested that they might constitute a separate genus ("an generis distincti"). This idea was readily adopted by Cassini (1827), who established the new genus Dorobaea for S. pimpinellifolius and its allies, without making any actual combinations, however. Bentham (Bentham & Hooker 1873) and later authors kept these taxa in the continuously over-expanding concept of Senecio, until the genus Dorobaea was revived by Nordenstam (1978). Jeffrey (1992) did not accept the genus as distinct from Senecio, but Bremer (1994) 32 Comp. Newsl. 27, 1995 followed Nordenstam in separating Dorobaea. Recent molecular studies confirm the distinctness of Dorobaea (Kadereit & Jeffrey, in press). Nordenstam in 1978 pro- visionally recognized one polymorphic species, D. pimpinellifolia (H.B.K.) B. Nord., citing the three other related species as synonyms, and mentioning S. callacallensis Cuatr. from Peru as probably belonging to the same alliance. We now present a slightly revised concept of species taxonomy in Dorobaea and make the formal transfers of S. /aciniatus and S. callacallensis to this genus. Dorobaea is a small Andean genus centered in Ecuador and with extensions into southern Colombia and northern Peru. Its species have a distinctive habit, being rosulate scapose herbs with conspicuous yellow or orange-coloured radiate solitary capitula. The disc-floret corolla is long-tubular, gradually widening towards the five- lobed apex, and the lobes are erect, narrowly ovate to lanceolate with a median resin duct, which may be more or less distinct. The style branches have a convex or shortly and obtusely conical tip surrounded by sweeping-hairs and sometimes provided with a short apical hair tuft, and the inner surface of the style branches have paired parallel stigmatic lines. The genus may comprise around five species, but at this stage we restrict ourselves to recognizing three species, which are keyed as follows. Key to the species of Dorobaea 1. Leaf-blade lanceolate to narrowly elliptic or lorate, lobes obtuse or rounded, often close or even overlapping; florets yellOW...........:.::cssesseseesesseecenseneeans sepeist ett Rie are. a, aes tee eatin ts 1. D. pimpinellifolia (H.B.K.) B. Nord. - Leaf-blade ovate or elliptic-oblong; florets orange or sometimes 2. Leaf-blade deeply lobed (to or almost to rachis), most lobes >1 cm long, distantly spaced from each Other: ..............ss0erssesniesseessest+otpadeten cones eens Te eee eee. Be. F 2. D. laciniata (H.B.K.) B. Nord, & Pruski - Leaf-blade shallowly lobed (to less than 1/2), lobes <1 cm long, CIOSEIVBDACEY, caiitetepaniteasei¥iasiaietidoutes 3. D. callacallensis (Cuatr.) B. Nord. & Pruski 1. Dorobaea pimpinellifolia (H.B.K.) B. Nord., Opera Bot. 44: 53 (1978). - Type: In herb. Humboldt & Bonpland (P). — Figs. 1,3 A-B. Senecio pimpinellifolius ("pimpinellaefolius") H.B.K., Nov. Gen. Sp. Pl. (folio ed.) 4: 136 , tab. 364 (1818). Comp. Newsl. 27, 1995 33 S. nubigenus H.B.K., loc. cit.; S. pimpinellifolius H.B.K. var. nubigenus (H.B.K.) Hieron., Bot. Jahrb. Syst. 28: 634 (1901). S. pediculariifolius ("pedicularifolius") H.B.K., Nov. Gen. Sp. Pl. (folio ed.) 4: 135 (1818). As recognized here D. pimpinellifolia is a quite variable species, especially in leaf shape and division. Also pubescence of involucre base and cypselas is variable. Usually the cypselas are shortly and appressedly white-hirsute (puberulous when immature), but some collections have quite glabrous cypselas. Phenotypic plasticity seems to be considerable, and plants are apparently strongly modified by altitude, exposition to light, and soil moisture. This species is widely distributed along the cordillera of Ecuador at elevations between 2300 and 4200 m, where it grows in cloud forest, scrubland and grass paramo, from dry soil to streamsides and bogs. It is also recorded from southern Colombia. 2. Dorobaea laciniata (H.B.K.) B. Nordenstam & J. Pruski, comb. nov. Basionym: Senecio laciniatus H.B.K., Nov. Gen. Sp. Pl. (folio ed.) 4: 137 (1818); S. pimpinellaefolius H.B.K. 8B laciniatus (H.B.K.) Wedd., Chlor. And. 1: 124 (1856); S. pimpinellifolius H.B.K. var. laciniatus (H.B.K.) Hieron., Bot. Jahrb. Syst. 28: 634 (1901). - Type: In herb. Humboldt & Bonpland (P, photo US! IDC microfiche 6209, card 105.II.3!). — Figs. 2, 3 C-D, 5. This species has a distinctive leaf shape, purple petiole bases and characteristically orange corollas. It has a more southerly distribution than the preceding species, in southern Ecuador (Loja Province), where it occurs at 2000 to 2500 m, on dry slopes. Two collections from Chachapoyas in northern Peru (Hutchison & Wright 4478, represented in 20 herbaria, King & Bishop 9203, US) also seem to belong here, although the rays are stated to be yellow. The southernmost record is A. Lépez 857 (US) from Peru, La Libertad, Otuzco, Agallpampa, 3100 m. Other Peruvian specimens from Cajamarca in the Province of Celendin (Hutchison & Wright 5211) also come Close, but differ by the densely villous and short petioles. The specimens from Cajamarca and La Libertad agree with D. laciniata, however, in the purple petiole bases and orange corollas. All specimens of D. laciniata examined have pubescent cypselas. 3. Dorobaea callacallensis (Cuatr.) B. Nordenstam & J. Pruski, comb. nov. Basionym: Senecio callacallensis Cuatr., Proc. Biol. Soc. Wash. 77: 151 (1964). - Type: Peru, Dep. Amazonas, Prov. Chachapoyas, middle E Calla-Calla slopes, near 34 Comp. Newsl. 27, 1995 Kms. 416-419 of Leimebamba—Balsas road, 2900-3100 m, 9 July 1962, Wurdack 1277 (US holotype! NY isotype!). — Figs. 3 E-F, 5. This species is easily recognized by the shallowly lobed, ovate leaf-blades. Petiole and scape are basally purple or lavender and the corollas are described as yellow in the type, and most other collections, but orange in one single collection less than 20 km from the type locality, viz. Boeke 1967, at km 399 of the same road (NY). The species is restricted to northern Peru. Further collections seen are: Peru, Dep. Amazonas, Prov. Chachapoyas, Cerros Calla Calla, 18 km above Leimebamba on rd. to Balsas, 3100 m, 7 June 1964, Hutchison & Wright 5587 (US); Dep. Amazonas, Prov. Chachapoyas, between Leymebamba and Calla-Calla, 2820 m, 27 May 1984, Smith & Cabanillas 7164 (MO, US); Cajamarca, Cutervo Prov., San Andrés de Cutervo, Parque Nacional de Cutervo, "Jalca" camino a la Laguna "El Pileo", 2680 m, 15 Mar. 1989, Diaz, Beltran & D'Achille 3310 (MO, US); Dep. Cajamarca, Cutervo Prov., Co. Hucan, Cutervo, 2850 m, 24 Jan. 1959, Velarde Nufiez 7147 (US); Dep. Amazonas, Prov. Pongard, Dist. Yambrasbamba, ca. 40 km N of Jumbilla, across R. Chiniaco from Yambrasbamba, 1860-2000 m, 2-26 Mar. 1967, S. S. Tillett 673-331 (US). Talamancalia H. Rob. & Cuatr. Another species with an anomalous position in Senecio is the Ecuadorian S. putcalen- sis Hieron., which we now refer to the genus Talamancalia H. Rob. & Cuatr. Senecio putcalensis matches Talamancalia by leaf-blades basally deeply lobed, by petiole bases commonly winged, by calyculate heads with many phyllaries, by orange corollas, by elongate and keeled anther apical appendages with somewhat thickened margins, by disc-floret corollas long-tubular with erect lobes, by similar style branch apices, and by ca. 8-ribbed pubescent cypselas without a strong carpopodium. This marks the first report of Talamancalia in South America. Talamancalia was recently described (Robinson & Cuatrecasas 1994) to accomodate two species from Costa Rica and Panama, respectively. The genus is close to the widespread neotropical genus Pseudogynoxys, which ranges from Mexico and the West Indies to Argentina with a concentration of species along the cordilleras of Peru, Ecuador and Colombia (Robinson & Cuatrecasas 1977). Pseudogynoxys comprises 14 species, which are vines or scrambling herbs, characterized inter alia by a pointed and penicillate tip to the style branches of disc-florets. The alternate petiolate leaves are generally entire with dentate to denticulate or entire margins, but a species with lobate leaves is being described from Brazil and Bolivia as P. lobata (Pruski in press). Another related genus is Garcibarrigoa Cuatr., a monotypic genus from northern Ecuador and southern Colombia (Cuatrecasas 1986). It is apparently close to Pseu- dogynoxys, differing mainly by the conspicuous, completely sheathing pseudostipules, Comp. Newsl. 27, 1995 and the non-scandent habit. Turner (1991) included Garcibarrigoa in Pseudogynoxys, but the genus was kept separate by Jeffrey (1992) and Bremer (1994). Robinson and Cuatrecasas (1994) again advocated its separation, and we share this view. Talamancalia differs from Pseudogynoxys by the non-scandent habit, winged petioles with basally pseudostipular appendages, the obtuse to rounded disc-floret Stylar tips with mainly lateral and apical, rather short sweeping-hairs, and the thickish and usually or perhaps always mucilaginous cypselar hairs. Talamancalia has some traits in common with Garcibarrigoa, including the non-scandent habit. Garcibarrigoa is very distinct, however, by its conspicuous and completely sheathing pseudostipules, and by its entire leaves with a strong and close venation and hirsute and dentate margins. In style morphology Garcibarrigoa comes closer to Pseudogynoxys than to Talamancalia, the style branch tips having longer and more pointed tufts of sweeping- hairs. A key to the three known species of Talamancalia is presented below. Key to the species of Talamancalia 1. Capitula 8-15 in each cyme; leaf-blades lobed only at very base ANAT Eee eee ns asbccssoocecvosvactboleens T. boquetensis (Standl.) H. Rob. & Cuatr. - Capitula 1-4; leaf-blades lobed in the lower half ........c.cccscscssssssssscsssscsssscesecescscecene 2 2. Leaves white-tomentose below; capitula 2-4 in terminal cymes on peduncles <1 cm long; calycular bracts ovate (Costa Rica) ....c.ccscssssesssssssssssssceseseeees EB 220005